Back in 1992, Hepburn raised the question of circannual rhythm and also! the idea that behavior within the colony is regulated by glandular secretions (such as queen substance, brood pheromone, etc.).

> The cohort involved in wax secretion and wax-working can now be added to a growing list of glandular-based and age-related polyethisms in honeybees. These results lead to the view that much of the flexibility in the division of behavior of honeybees operates within constraints imposed by underlying physiological cycles of glandular activity. The real factors, i.e. climate (photoperiod, temperature etc.) or a basic endogenous rhythm, regulating the circannual rhythm of a colony need to be established. 

> ... there is now sufficient information to ask how the probability of a cohort performing given tasks is constrained by underlying physiological characteristics (glandular secretions) and whether the tasks are susceptible to modulating stimuli (queen pheromones, nectar influx etc.).

Muller, W. J., & Hepburn, H. R. (1992). Temporal and spatial patterns of wax secretion and related behavior in the division of labour of the honeybee (Apis mellifera capensis). Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology, 171(1), 111-115.

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